Decadent Dinosaurs: Directed Evolution in British and North American Literature, 1890s-1970s. (2024)

In her 1951 book A Land, British prehistorian Jacquetta Hawkes reflected on a "merciless force in evolution" that caused "trends, once they have begun, to become excessive and at last pathological" (29). Her example was the fork-horned Miocene deer Synthetoceras (fig. 1), which "must have looked more ridiculous than Munchausen's stag with a cherry tree sprouting from its forehead; it is not surprising that it found life intolerable and rapidly became extinct." (1) While Baron Munchausen's claim to have produced a tree-horned stag by shooting it with cherry stones could be dismissed as fantastical, nature itself, Hawkes proposed, could produce comparable monstrosities. Her understanding of the "ridiculous" horn drew on two influential paleontological concepts: orthogenesis (evolution along a linear track) and phylogeronty (the parallel between the lifespan of an animal group and the lifespan of an aging individual). In short, that is, Synthetoceras's aberrant horn, an overly specialized structure unfitted to its environment, was a sign of the genus's evolutionary old age. Other groups purportedly trapped in evolutionary senescence included "luxurious forms of life" (79) like the armored dinosaur Stegosaurus and the "bizarre and decadent" (81) ammonites.

This article explores the heyday of these suggestive concepts in British and North American science writing and popular fiction. To this end I will group orthogenesis and phylogeronty, along with recapitulation (the belief that growing animal embryos rapidly reenact their own evolutionary histories), under the umbrella term "directed evolution." In this I follow Warren D. Allmon (2020: 424), who carefully distinguishes "patterns in the history of life" from "views of the processes of evolution." While the identification of the former is a descriptive affair, the identification of the latter, with which this article is concerned, is explanatory. Allmon observes that, for many paleontologists who examined trends like the evolution of increasingly spectacular horns, "recognition of the seemingly ubiquitous pattern frequently slid without comment into acceptance of the process" (441). As this slippage suggests, directed evolutionary hypotheses leaned heavily on metaphor and analogy, chiefly analogies involving the life cycle: youth, maturity, senility. Orthogeneticist geologist Herbert Leader Hawkins remarked in 1920, for example, that "the analogy is too perfect to be the outcome of mere coincidence or sophistry; it is an expression of the simplicity and order that are fundamental attributes of the Universal Cosmos" (118). Similarly, paleontologists compared the inertial evolution of nonadaptive traits to the decadence of the Roman Empire (Swinton 1934: 179), the screeching brakes on the London Underground (1966: 117), and even the disconsolate final movement of Pyotr Ilyich Tchaikovsky's Symphony No. 6 (Hawkins 1920: 117).

Charles Darwin famously struggled to portray natural selection as nonteleological (Beer 2009: 76), but, for naturalists skeptical about the importance of selectionism in generating species, teleological or at least directional analogies were critical. Devin Griffiths (2016: 18) distinguishes "formal analogies" that "apply a previously understood pattern of relationships to a new context" from "harmonic analogies" that "allow significant shared features to emerge." Unlike the bilateral harmonic analogies Griffiths sees as central to Darwin's insights on the origin of species, such as the comparison between domestic and wild breeding (34), the analogy between the individual life cycle and that of the animal group was a formal superimposition of the former onto the latter, explaining evolutionary trends through a ready-made model. As already indicated by the evocation of Rome, this ostensibly naturalistic analogy often involved a cultural value judgment, comparing waning genera to decadent civilizations. The notion that civilizations rise and decay in cycles is usually considered a figurative understanding of human cultures deriving from the literal cycles of nature. Against this grain, Whitney Davis (2005: 138) contends that "decadence is really a metaphor of Culture applied to Nature." This is the case in the sharpest writing on directed evolution, according to which the persistent maladaptation of animal groups indicated an unconscious but quasi-moral failure.

Directed evolution, which was gradually excluded from mainstream anglophone paleontology in the decades following the 1940s, has long been the subject of scholarship by historians of science (Bowler 1989; Manias 2017). Scholars of twentieth-century literature have paid it less attention, despite an increase in literary work on non-Darwinian evolutionary theory. Intense interest in literature and Darwinism (Greenberg 2009) has been accompanied by literary research on theories that downplayed the importance of natural and sexual selection like neo-Lamarckism (Hale 2006), Hugo de Vries's mutation theory (Endersby 2013), and Henri Bergson's vitalism (Moses 2014), but paleontology and especially directed evolution have usually been only passing subjects of analysis in this work. Cathryn Setz (2019: 62-75) does briefly explore James Joyce's irreverent treatment of orthogenesis in early versions of Finnegans Wake (1939), and several scholars have examined Virginia Woolf's interest in evolutionary specialization and embryonic recapitulation (Chan 2014: 174; Livingstone 2022: 151-95). I will return to Woolf, but for now it is enough to note that avant-garde modernists like Joyce and Woolf usually opposed the deterministic assumptions underlying directed evolutionism. In contrast, the figures I examine here were typically drawn to its potential relevance for halting societal decline, and they relied on more traditional literary genres.

This ideological and stylistic conservatism may help illuminate why the currency of decadence in paleontological thought has not found a significant place in the growing body of work on literary decadence in the twentieth century (Hext and Murray 2019) or in relevant research on decadent ecology (Denisoff 2021) and decadent science more generally (Kistler 2019). Where the decadent movement is broadly associated with the celebration of aestheticism, urban life, individual expression, strange sympathies, and provocative gender politics, scientists discussing decadence tended to be less enthusiastic about these associations. Evolutionary thinkers regularly warned about the longterm dangers of becoming hereditarily unfit for one's environment, but theorists of directed evolution, for whom unfitness was potentially pre-programmed, had particular cause for vigilance. The most vocal called for humans to avoid the indulgent overspecialization that had, they thought, led animals to evolve in a spectacular but self-destructive manner, a danger for which they sometimes proposed eugenic solutions (Rieppel 2019: 163-65). In focusing on directed evolution, I will thus consider sources mostly other than classic studies of the related literature of degeneration (Greenslade 1994), which have drawn on psychological and medical more often than on paleontological discourses. While naturalist E. Ray Lankester, in 1879, famously cited the fall of Rome when contending that excessive comfort led to decreased anatomical complexity in sea squirts (Bowler 1989: 335-36), the bizarre fossil discoveries of the fin de siecle and beyond seemed to suggest that, at least for nonhuman animals, decadence was inevitable. It was more often indicated by heights of complex overspecialization, evidenced in bizarre ornamentation, than it was by the degenerate neoteny of the sea squirt.

Directed evolution was theorized and explored in writings for specialists and general readerships in the anglophone sphere for over half a century, and the fluidity with which pronouncements on dinosaurs moved to reflections on the fall of empires makes it richly interesting for literary scholars. I will, after a brief introduction to its scientific history, focus on three representative manifestations of directed evolution in texts that have mostly received little critical attention. First, I examine popular romances by authors like Weird Tales stalwart Robert E. Howard, in which dinosaurs and decadent civilizations both function as foils to potent protagonists. Second, I compare H. G. Wells's Mr. Blettsworthy on Rampole Island and Marie Stopes's Love's Creation, two novels published in 1928 in which the apparently deterministic and directional nature of evolution is implicated in contrasting social reform agendas. Finally, I demonstrate how prominent scientists relied on theatrical metaphors to temper the otherwise pejorative characterizations of phylogerontic animals with a tragic pathos. In sum, I argue that directed evolution gave a culturally recognizable shape to prehistoric and modern life alike, figuring extinct animals as embodying moral lessons about the route to human evolutionary success. This had a bracing appeal to elite and middle-class reformers who hoped that the most industrious humans could break out of the grooves condemning nonhuman animals to extinction. But such optimism contended with an unease about the ultimate failure of humanity and a sense that, in contrast with the spectacular features that emerged in decadent genera like Synthetoceras, evolutionary health required mediocrity.

Senile or Decadent

Directional or cyclical interpretations of life's development have a long history, but in the late nineteenth century, especially in Germany and the United States, these ideas were formulated in evolutionary terms. To cite two crucial contributors from a wide pool of thinkers (Popov 2018: 7-28), German biologist Ernst Haeckel's epochal work on the evolutionary recapitulation performed by embryos was complemented, in the United States, by Alpheus Hyatt's research into ammonite shells, the increasingly bizarre nature of which, he contended, foreshadowed the extinction of the group (Allmon 2020: 441-43). "Orthogenesis" (428) was coined in 1893, as was "phylogerontic," although the latter clinical Greek coinage was often ignored in favor of the more ominous terms "racial senescence" or "racial senility" (442). From the turn of the century on, the Yale Peabody Museum and the American Museum of Natural History in New York became centers of directed evolutionary thought, while at the Natural History Museum in London paleontology was similarly shaped by orthogeneticists like Arthur Smith Woodward, William Dickson Lang, and William Elgin Swinton.

As it emerged in the nineteenth century, orthogenesis overlapped in some respects with theistic interpretations of life's progressive unfolding. Evolutionary skeptics like Canadian evangelical geologist John William Dawson shared with avowedly agnostic orthogeneticists an organizing model of rise and fall (1873: 182-83), rejecting, in Peter Bowler's (1989: 348) words, "evolution governed by 'chance.'" Paleontology's already precarious relationship with natural theology, however, was challenged by the evidence that animals could develop detrimental physical features. In the early nineteenth century, knowledge of the monstrous hybridity of many extinct animals was often accompanied by Paleyan arguments that their design was divinely ordained and thus functional. The South American giant ground sloth Megatherium, a Victorian byword for ponderous obsolescence, was characterized by Anglican paleontologists as perfectly fitted to its environment (Dawson 2011: 207). Although orthogenesis could be seen as adaptively benevolent during periods of evolutionary youth and maturity, as in the case of the fine-tuning of the horse into a paragon of speed and elegance (Osborn 1917: 267) or the mammal brain's expansion, it left countless extinctions in its wake, including not just the ammonites and armored dinosaurs but also Megaloceros, another deer known for its strange antlers, and the saber-toothed cats, with canines that appeared detrimental to predation. Adaptive explanations for these dazzling structures were not widely persuasive.

The idea that evolution was internally and linearly directed, which appeared in many variations, was one among several competing theories, into some of which it occasionally bled. Thriving into the 1930s among paleontologists, directed evolution was ultimately overshadowed in anglophone science by the "modern synthesis" of natural selectionism and population genetics (Cain 2013) (although orthogenesis itself survived into and beyond the Cold War years [Popov 2018: 73-112]). It provoked robust objections not only from specialists, as demonstrated in a letter from Alexander Hamilton Gunner (1935) of the nonprofessional Geologists' Association to William Elgin Swinton at London's Natural History Museum that chastised orthogeneticists for their "loose thinking," insisting that the "sabretooth of course could feed and fight" and that "each creature succeeded within its own time." Swinton, however, held that directed trends caused anatomically generalized (and thus evolutionarily flexible) animals to overspecialize, to grow to unsustainable sizes or develop cumbersome excrescences. At Yale, Charles Emerson Beecher (1898: 355) represented one such development in a chart, mapping ontogenetic (individual) age onto phylogenetic (evolutionary) age to show how both tended toward spinosity (fig. 2). In discussing such "unwieldy" animals as barely fit to live, paleontological writers adopted a "grotesque" mode, comparing them to gothic ornamentation or the nonsensical creations of Lewis Carroll (Fallon 2021: 63-98). In one typical aside, Swinton (1948: 216-17), perhaps Britain's most famous paleontologist in the mid-twentieth century, joked that "mighty dinosaurs" like the "grotesque" bone-headed Pachycephalosaurus "were a mighty nuisance to themselves." Half a century prior, in the American Journal of Science, Beecher (1898: 354) held that, just as the "decadence" of an elderly animal "produces extravagant vagaries of spines," evolutionarily senescent lineages produce "extravagant types" like the three-horned dinosaur Triceratops. In light of the era's expanding imperialism, Beecher's framing of inadaptive traits as "decadence" reflects the Western fascination with the legendary luxuriance and resultant destruction of the Roman Empire (Malik 2019). Uneasy reflections on the causes--or perhaps inevitability--of the fall of empires permeated nineteenth-century culture (Buckley 1966: 66-93) and Beecher's theories offer just one instance of their scientific expression. In another, the critic John Addington Symonds (1890, 1:46) evoked evolution and "organic growth" to posit the "natural end" of genres in art and literature. Fin de siecle aestheticism celebrated decadence's potential for cultural transformation and disruption (Gluck 2014: 355; DenisofF 2021: 32) and many of the features conservative critics denounced, and aesthetes like Oscar Wilde celebrated--excess, effeminacy, delicacy, disproportion, inefficiency, ornamentation, futility--were precisely those of the "grotesque bizarrerie" (Lull 1917: 518) paleontologists mapped onto doomed extinct animals in the following decades. While in the 1890s the decadent avant-garde did not embrace paleontological parallels, they were exploited by reactionaries like the Marquess of Queensbury, who mockingly compared Wilde to the Cretaceous dinosaur Iguanodon (Greenslade 1994: 70-71).

The process of tying together the declining adaptability of animal groups and the decadence of empires continued in the twentieth century. In The Dinosaurs (1934), an influential text addressed to specialists and educated general readers alike, Swinton wrote that "senile or decadent" (181) as the last dinosaurs were, they represented "a mighty nation whose glorious history and empire have crumbled to the dust" (208)--implying that humans, or perhaps nations or races, ought to learn from the dinosaurs' failure. The significance of directed evolutionary trends for human culture was considered by thinkers as divergent as Oswald Spengler (1926-28, 2:32) and Rabindranath Tagore (1931: 28-29), so that political understandings of such trends were by no means uniform, beyond their appeal to grand narratives. Hyatt argued that granting women's rights would accelerate human extinction (Bowler 1989: 341); for American educator Granville Stanley Hall (1904, 2:561) it was men who were "a trifle senile, if not decadent"; in his Glimpses of the Ages (1905-8, vol. 1), Jamaican-born doctor Theophilus Scholes correlates the geological "law of progression" (387), by which mammals overtook reptiles, with a cyclical model of empires, concluding that European supremacy might be overtaken, in the "next cycle" (390), by "Ethiopic" (392) civilization.

It is unsurprising that directed evolutionary theories were applied to human society, and understood in racial terms, given that "racial senescence" was usually the designation preferred over "phylogeronty." The slippages facilitated by this terminology recall those that followed Darwin's use of the polyvalent word "race" in On the Origin of Species (1859) (Beer 2009: 47). Geologist Innokenty Pavlovich Tolmachoff (1928: 1137), for example, speculated that the lack of adaptive flexibility in many "extinct animals" resulted from "lost racial vitality." Similarly, British eugenicists argued that white unfitness would have severe consequences for the Empire (Bland and Hall 2010: 213), and though in general directed evolution wasn't an attractive framework for the eugenics lobby--whose utopian vision didn't accord with a belief in the decline of species (Saleeby [1909]: 99-101; Bather 1920: 86)--some felt it might be harnessed and redirected in order to avoid decline. For example, as doyen of the American Museum of Natural History, the orthogeneticist Henry Fairfield Osborn presided over exhibitions that espoused evolutionary progress, and, with his elite clique, believed that promoting immigration restrictions and eugenics could eliminate the unfit and non-Nordic from the American gene pool (Rieppel 2019: 144-45). Directed evolutionism was thus as often a spur to action as it was a cause for fatalism.

Close to the Primitive

Authors writing in the lost world romance genre posed their own solutions to decadence. For Bradley Deane (2014), "lost-world romancers" (148) like H. Rider Haggard ploughed their heroes through the wilderness, seeking in Indigenous peoples a "primal strength" that had "eroded in the degenerate metropole" (149), a process that was reworked in the American pulps. On both sides of the Atlantic (Noble 2016: 54-57), moreover, these ultramasculine narratives came to also feature ferocious extinct animals. Linking such work to Osborn's social policies, Marianne Sommer (2007: 319) suggests that in romances like Edgar Rice Burroughs's The Land That Time Forgot (1918), the primordial lost world setting acts as an "incubator of evolution" for white male protagonists. In such works heroes are tested against powerful Indigenous peoples and the wild prehistoric animals they're associated with, symbols of vigor regularly juxtaposed with the corrupted inhabitants of decaying civilizations. This juxtaposition reflects interest not just in degeneration but also in orthogenesis. Dinosaurs represented raw and "primitive" nature, but also inadaptive decadence, and in this we can read them as objective correlatives: they embody the dilemmas facing the protagonists, who must always defeat both physically terrifying monsters and morally depraved human (or humanoid) foes, drawing both on brute force and on civilized urbane cunning. Running through these stories of men taking control of the evolutionary steering wheel, we thus find a persistently orthogenetic aesthetic of bristling reptilian survivals and gilded but rotted citadels.

The conventions of this subgenre of lost world romance began to congeal following C. J. Cutcliffe Hyne's The Lost Continent, set in prehistoric Atlantis and serialized in British and American editions of Pearson's Magazine from 1899 to 1900. In Hyne's story, we find this ancient transatlantic civilization in its Wildean decadence: "posturing" and "mincing" (1900: 91), Atlantean men attend only to superficialities while their ascetic priests retreat from society; all are ruled over by an evil femme fatale empress named Phorenice. The story is set in a nebulous time, when humans share the planet with giant prehistoric animals: Phorenice rides a mammoth, plesiosaurs haunt the waves, and dinosaurs stalk the frontiers. The protagonist, Deucalion, an official of Atlantis's Mesoamerican provinces, is repulsed by the effete Atlantean metropole. He exiles himself in disgust to the frontier, toughening himself among "colossal lizards" (274) like the Brontosaurus--a herbivorous dinosaur here depicted as a predator covered, notably, in "horny plates" and "spines" (275)--before returning to decadent Atlantis in an effort to reverse its decline.

Burroughs incorporates similar elements. In Tarzan the Terrible, a novel serialized in Argosy All-Story Weekly in 1921, the lost valley of Pal-ul-don is inhabited by a predatory animal called the gryf, that Tarzan, who has "seen its skeleton in the museum in London" (65), identifies as a descendent of Triceratops. Burroughs was familiar with teleological interpretations of evolution (Reid 2018: 216-26), and likely knew that Triceratops was considered the last of the dinosaurs, its horned head a textbook example of how racial senescence turned useful structures into hindrances. Exhibiting even more spinosity than Beecher diagnosed in its ancestor, the "grotesquely serrated" (10) gryf has "three parallel lines of bony protuberances down the back" and the blunt "hoofs" of Triceratops have evolved into "talons" (65-66). The bestial Tarzan, however, admires the "courage and strength" of the dinosaur, which he uses as a steed. In the course of the novel Tarzan encounters two distinct groups of racialized ape-men (both evolutionarily stunted, rather than decadent, although the possibility of atavism is raised): the nature-loving black Waz-don and the white, urban Ho-don. Tarzan's clash, significantly, is with a portion of the latter race, the effete and hypocritical priests who have corrupted the grand Ho-don city of A-lur with their cruel and extravagant rituals. Though Burroughs plays fast and loose with the established paleontological symbolism, turning a decadent dinosaur into a symbol of vigor, his evolutionary intervention is nonetheless clear: with the power of evolutionary plasticity, the gryf-riding Tarzan defeats the priests and reverses the decline of the debauched Ho-don metropolis--unsettling the linear rails leading toward senescence.

Sharing many elements with Burroughs's story, his Texan contemporary Robert E. Howard's Red Nails (1936), which was serialized in Weird Tales, bears many similarities to Tarzan's adventure. At its start, Conan the Barbarian is attacked by a "dragon" covered in "serrated spikes," a "monstrous survival of an elder age" (Howard 1936a: 21-22). Although Conan, like Deucalion and Tarzan, is "too close to the primitive himself" (25) not to feel some "kinship" with the ancient monster, he poisons it, deploying a combination of manly vigor and quick thinking. This act distinguishes him from the decadent inhabitants of nearby Xuchotl, who believe that "no man ever killed a dragon" (Howard 1936b: 213). This city is shut off from the outside world; locked in the city and constantly murdering each other, the inhabitants of Xuchotl are propelling themselves toward extinction. But these are not the original inhabitants. The opulent city's original population was an eastern race who used "necromantic arts" to revive the "dragons ... whose monstrous bones they found in the forest." Predictably, "luxurious sloth" (214) rendered the original dwellers of Xuchotl easy prey to barbarian invaders, the undead saurians gradually dying off (again), with Conan slaying the last survivor. Another adaptable hero capable of dispatching both destructive dinosaurs and senile urban cultures, Conan eventually purges the debased Xuchotl of all its inhabitants. The lurid phylogerontic vision driving this story reflected Howard's exchanges with his friend H. P. Lovecraft. Lovecraft's letters brooded on Spengler's diagnosis of Western "cultural senility" (Lovecraft and Howard 2017: 1:246) and lectured Howard on "what Haeckel called the first and basic law of all biogeny," asserting that "the history of any life-series is repeated in the development of the individual from egg to senility" (2:748).

In another similar novel, the role of adaptable hero was actually filled by a dinosaur. Scottish-born American mathematician E. T. Bell's Before the Dawn (1934), published under the name John Taine, depicts an invention by which fossils can project their own life histories as three-dimensional movies. In the lives of dinosaurs we thus see processes of directed evolution brought into relief when the projector speeds up the fossil film, so that the rise and decline of genera appears "no more startling or unnatural than the slow growth to maturity and the gradual decline to old age and death of a single human being" (Taine 1934: 131). The audience of scientific spectators watches as dinosaurs' once useful "horny armor ... continued to develop like a horrible disease," resulting in "starving senility" (142), one observer expressing shock that nature could produce such "a meaningless, helpless monstrosity" as the long-necked sauropod dinosaur dubbed the "lotus eater" (69). Persistently, the novel associates dinosaurs with Orientalist images of decadent luxury and despotism, giving specimens biblical names like Belshazzar, Satan, and Jezebel. (2)

In a surprising twist, the tyrannosaur Belshazzar, like the hero of a pulp romance, begins to master this decadent environment. Predatory dinosaurs were not usually understood as having evolved in the phylogerontic direction that herbivores like the sauropods, ceratopsians, and stegosaurs had, so that their status in the directed evolutionary narrative appeared uncertain--perhaps helping explain why the vigorous and thus faintly admirable dinosaurs in the lost world romances I have discussed were all represented as predatory (even if, in the case of Hyne's Brontosaurus and Burroughs's Triceratops, this meant getting the science wrong). Bell's narrator recognizes the predatory "bipedal lizards" as a "great race," indeed, as "one of nature's major efforts" (120-21), and Belshazzar himself is described as "a perfectly balanced engine of aggressive destruction" (160) (words echoing Osborn's [1917: 214] memorable description of Tyrannosaurus rex as "the most destructive life engine which has ever evolved"). But Belshazzar appears somewhat sympathetic not simply for his physical power; he is depicted also as one of "the pioneers of science," his methods of survival revealing knowledge of "the laws of levers," of the "law of falling bodies," and of the principles of "hydrostatics" (219-21). When he is killed, the novel ends with Belshazzar's "last snarl of defiance" (247). In all these stories, a laudable protagonist is defined by a capacity to adapt to shifting environments, to draw from whatever is demanded to contend with the seemingly inexorable processes of decay and extinction.

Survival of the Rotten

The directed evolutionary ethics loosely informing pulp romance and science fiction could be laid out more prescriptively in novels of ideas like Marie Stopes's Love's Creation (1928). Although by the time of the novel's publication Stopes was best known as a birth control campaigner, her previous career had been that of a professional paleobotanist, and she worked at the University of Manchester from 1904 to 1910. Stopes was versed in the various evolutionary mechanisms then in circulation and may well have discussed them with D. M. S. Watson, a student with whom she produced an influential paper on coal (Chaloner 2005: 132), and who went on to argue persuasively in favor of orthogenesis in amphibians (Bowler 1996: 238-39). Carla Hustak (2014) has pointed out some continuities between Stopes's paleobotanical thought and sexological thought, but Love's Creation stands as one of her frankest applications of paleontological research to twentieth-century social life.

Stopes's 1928 novel concerns a central romantic pair, University of London biologist Kenneth Harvey and progressive reformer Amber Rose. After his first wife dies, Kenneth wanders about in South and Southeast Asia, where what begins as racist revulsion at Asian populations spurs him to search for the "bridge" (227) linking him with the Other; he turns to Buddhist and Daoist texts for insights about the individual's relation to the whole, and he turns to the study of paleontology (302-3). Finally, he arrives at his "big illuminating idea" (264), which is explained at length in a chapter prefaced with the forbidding warning that it "should only be read by those who think" (317). Kenneth has been persuaded by "the bigger Palaeontologists, like Osborne [sic] and Smith Woodward" that, contra the belief that natural selection promotes superiority, animal groups "die out automatically," each "following a definite course of procedure more or less regardless of their environment" (325-27). Kenneth believes that his variant of orthogenesis will be paradigm shattering: individual lives, he argues, are mere components of a "larger unit," much as "individual cells composing our brains" produce "a super-something--a consciousness" (328-29), and he compares the way extinct species within this larger unit have been "jerked off the rails" to the metamorphosis of a "caterpillar" (330). Phylogeronty and extinction, then, represent the aggregate consciousness sloughing otf elements not needed for a higher evolution.

In the words of her biographer, Stopes "was an elitist, an idealist, interested in creating a society in which only the best and the beautiful should survive" (Rose 1992: 134). There was thus a "strong eugenic strain in her views," Bland and Hall (2010: 217) note, albeit one of a rather "maverick" character, and as Stopes's warning away all but "those who think" suggests, her eugenic elitism manifests in Kenneth's big idea. "Diseased city dwellers" should be thought of not as "isolated strains doomed to individual extinction," he argues, but as "festering sores" in the "Greater Consciousness" (340-41), and it is "our responsibility" to "destroy the diseased fragments" (341-42). Like Osborn, whose work Kenneth approvingly cites, Stopes's author-surrogate here suggests that, using eugenic methods, the orthogenetic cycles of evolution can be harnessed for the health of the human species, which, given the tenor of her pronouncements (Rose 1992: 147, 155), seems to mean white middle-class members of the species. Supplementing the negative eugenics Kenneth more or less proposes, he and Amber practice something like positive eugenics: after his first wife's death, Kenneth's mother laments "that her son's life would no longer develop along the lines she anticipated" (182), but his more sexually compatible marriage with Rose reveals that this initial derailing, to mix metaphors, has released both lovers from their "chrysalis" (412).

It's illuminating to contrast Love's Creation with another novel published in 1928, H. G. Wells's Mr. Blettsworthy on Rampole Island. The titular protagonist finds himself stranded on an island of cannibals, who themselves are terrorized by villainous specimens of the prehistoric ground sloth Megatherium, "grotesque monsters" (142) who survive by obstructing the progress of other lifeforms. Though Wells was no committed orthogeneticist, and previous studies of Blettsworthy (Philmus 1998: 32-40, e.g.) have not attended to directed evolution, I suggest that it nonetheless plays an important role in this novel. It's true that he had adopted a chiefly Darwinian framework in his first evolutionary epic serial, The Outline of History (1919-20). The Science of Life (1931 [1929-30]), a later work coauthored by Wells, his son, and biologist Julian Huxley, even threw "the shadow of doubt" (350) on directed evolution in favor of selectionist arguments and considered the notion of "racial senescence" merely a "loose" analogy, attributing extinction not to "inner ageing of the germ-plasm" but to "changes in outer conditions" disadvantageous to species that had "moulded themselves too perfectly to the passing world" (382-83). At least one author, however, found Wells's novel an apt representative of directed evolutionary thought: in The Dinosaur in East Africa (1930), geologist John Parkinson introduced his view of the dinosaurs' "degenerate senility" (173) with an epigraph drawn from Blettwsorthy's observation that, "Far from Evolution being necessarily a strenuous upward progress to more life and yet more life, it might become, it could and did evidently in this case become, a graceless drift towards a dead end" (Wells 1928: 171-72).

Indeed, the notion of phylogeronty permeates Wells's novel. Before becoming stranded on Rampole Island, Blettsworthy had believed that a species facing changed conditions "produced an adaptation to fit it beautifully to these new conditions." Observing the island's repulsive ground sloths, however, he finds that "species under altered conditions did the queerest and most futile things" (170) and Blettsworthy concludes that "animals do not necessarily survive by being swifter, stronger or wiser," that the "survival of the rotten and dying is possible" (171). The choice of Megatheria was pointed. Wells likely knew that the extinction of this animal was not conventionally attributed to phylogerontic maladaptation. Rather, he was sarcastically bringing into a twentieth-century context the nineteenth-century natural-theological arguments I mentioned above, according to which Megatheria exemplified how a superficially monstrous animal was perfectly fitted for survival--a continuation of the assault on theistic Victorian science initiated in the novel's early scenes, where Blettsworthy's thinking is shaped by his naive uncle, a fusion of Pangloss and Charles Kingsley, who repeats that "all things work together for good" (32). Indeed, by the early twentieth century, mid-Victorian thinkers were coming to be seen as "Megatheria" themselves (Smith 1918: 230).

Outmoded Victorian science was not Wells's primary target, however. Later in the novel, Wells reveals that Blettsworthy's adventures on Rampole Island were a trauma-induced hallucination--that the Megatheria were his mind's symbolic projection of the failing liberal institutions of Western Europe. And it is only during the Great War, the catastrophic result of these failures, that Blettsworthy regains full consciousness. Presumably, the novel's quasi-allegorical form exemplified the superior "technique" for a "novel of ideas" (Rose 1992: 189) Wells alluded to in critiquing Stopes for the metaleptic framing of her chapter on orthogenesis. In contrast to Stopes, Wells wished to avoid interrupting the reader's suspension of disbelief, the novel's critique emerging only gradually: even before he regains lucidity, in Blettsworthy's mind "Institutions had become entirely confused with Megatheria and Megatheria with Institutions" (174). He accepts the idea that the "institutions of mankind came just as much within the scope of biological generalization as the life of any other living being," so that, just as Rampole's Megatheria obstruct youthful species, "obstinate obstructives" (172) linger in human society. After the War, a despondent Blettsworthy is invited to the National Liberal Club by Lyulph Graves, an old acquaintance who, like the socialist Wells himself (Toye 2008: 184), retains faith in aspects of liberalism, even as he thinks that the megatherial Liberal Club represents "a banner of progress that someone has forgotten to carry on" (279) since its "Gladstonian" glory days in the nineteenth century. Moribund Victorian organizations, however, "can be changed without any change of nature" (287). By maximizing the "efficiency of the educational machinery of mankind," Graves contends, "Megatheria can die" (285). On this view, political rationalization is possible, but not without serious educational reform.

The novel's skepticism about the inevitability of evolutionary and political progress reflected a career spent countering complacent views like those held by Blettsworthy's uncle and, initially, by Blettsworthy himself. Adam Roberts (2019: 347) notes Mr. Blettsworthy on Rampole Island's position in a "feedback loop" running back to Wells's rejection of evolutionary optimism in The Time Machine (1895), which, Bowler (1989: 337) observes, pointedly depicts evolution as nondirected: the murderous Morlocks and feeble Eloi of the far future represent degenerate adaptations by different social classes of humans to their environments. Wells's (1896: 595) contemporaneous journalism rejected the "clamour for the Systematic Massacre of the Unfit" and insisted on the all-importance of education to avoid a decadent future. Although he consistently prioritized education, as shown in Blettsworthy, the technocratic Wells, like Stopes, also came to endorse negative eugenic policies, particularly regarding the suppression of certain mental illnesses and disabilities (Stover 1990). Unlike Stopes, however, he saw eugenics as a less reliable tool for creating an ideal society than socialist mass education (Partington 2003: 77).

With the "old-young Megatherium" (279) in Blettsworthy, Wells suggested that, while directed evolution might not operate in the natural world, elements of it might be at play in the political world. Surviving past the end of their life cycle even though they are not fit and not breeding, the sloth institutions comically defy selectionism. Most proponents of orthogenesis suggested that while phylogerontic genera faced extinction when ecological conditions changed, under steady, amenable conditions, overspecialized dinosaurs like Triceratops might endure. The Megatheria of Rampole Island thus achieve endless old age by preventing changes in their habitat (parodically reversing Wells's precept, noted by Simon J. James [2012: 126-27], that static societies face "extinction"). In stressing that the senile Megatheria represent political institutions, then, Wells denaturalizes the evolutionary logic of his story. Where Love's Creation would have society lean into the inevitable direction of evolution by pruning phylogerontic individuals, Blettsworthy preaches amelioration by changing the environment, eliminating senescent political structures by means of left-wing educational reforms. Throughout the novel, as Blettsworthy grows disillusioned with the liberal assumptions of his youth, he feels his body, in phylogerontic manner, is "in hideous discord with the entirely inhospitable world into which it had come" (88), but the problem isn't in his body itself but in the inhospitable world produced by the unreformed Victorian social structures. Graves thinks that, rather than forcing the inadequate individual to fit the environment or die, the inadequate environment may be adapted for the good of the individual, and, whether or not we are intended to agree with a character who once swindled Blettsworthy out of his money, Wells gives us little alternative.

During the Second World War, his interbellum hopes for educational meliorism thwarted, an embittered Wells returned to directed evolutionary language. In his Nature article "The Illusion of Personality," for example, he was no longer dismissive of phylogeronty, citing "repeated" paleontological precedent for the emergence of a "flurry of abnormalities before the collapse and obliteration of some dominant group which has outstayed its welcome" (Wells 1944: 395). Now he judged that the same phylogerontic pathology that produced "abnormalities" (395) in extinct genera "seems to be the case with man to-day," given that "man" offers "no collective resistance in the face of change" and "persists in his follies" (397).

Fragments of Old Playbills

For Blettsworthy's friend Graves, the megatherial vestiges of Victorian liberalism do not "know how to get off the stage" (279). And indeed, the notion of prehistory or, for that matter, history, as a stage was a venerable one (O'Connor 2007: 377-84). Both animal groups and geological eras, much like individual lives and civilizations, were framed as reaching a "climax" followed by a denouement and curtain (Osborn 1917: 183, 210, 225), and Shakespeare allusions, especially evolutionary spins on Jaques's outline of the seven ages of man in As You Like It (Hutchinson 1889: 260-61), seemed to come instinctively to the pens of British paleontologists (Parkinson 1930: 56, 91, 103). "Shakespeare has made us familiar with human growth-stages," casually noted William Dickson Lang (1919: 102) in a paper on "Old Age and Extinction in Fossils." Also alluding to Jacques's vision, Francis Arthur Bather (1928: lxxxiv), Keeper of Geology at the Natural History Museum, observed that "an eventful history" would postpone an evolutionary "second childhood." Of course Shakespeare allusions are ubiquitous in modern British writing, but their persistent association with directed evolutionism in particular nonetheless points clearly to how writers on evolution found in Shakespeare the same tensions between inevitable decline and individual agency that characterized directed evolutionism. Audiences can identify Macbeth's hamartia, but they cannot prevent his fall. Harmful orthogenetic trends can be identified, but can they be halted?

Viewing directed evolution through a Shakespearean lens was natural for Bather (1887: 83), a paleontologist and also author of scholarly works discussing "the physiology, morphology, and embryology" of "the puns of Shakespeare." Having contributed to orthogenetic vocabulary at the fin de siecle, by the 1920s Bather was experiencing doubts. He warned the British Association that, while the idea that genera let out a "despairing outburst before death" was "delightful as metaphor," it likely stemmed from "ignorance alone" (Bather 1920: 85), though these doubts did not always temper his sense that extinction was as gloomily preordained as the fate of characters in a Shakespearean tragedy. In the peroration of a 1928 address to the Geological Society, Bather (1928: xcvi) reflected that countless "Classes and Orders" had finally lost the "flexibility ... with which to meet new conditions," putting on "a short-lived majesty" and fading before even learning "to wield the sceptre and to support the crown." Bringing humanity into this narrative, Bather imagined an Anthropocene wasteland of the future in which "Man" is forced, "more literally than Hamlet, to 'feed on air,'" as the human races decline toward "the inexorable end." With these words he launched into Hotspur's dying lines in Henry IV, Part 1:

 But Thought's the slave of Life, and Life, Time's fool; And Time, that takes survey of all the world, Must have a stop. (xcvii)

Bather, a specialized invertebrate paleontologist, wryly observed that he was aptly ceding his presidential title to an "unspecialized" geological generalist, J. W. Gregory. Inelastic species, kings, and presidents of the Geological Society alike faced inevitable obsolescence.

Bather's dramatic flair reflected a more empathetic approach toward these evolutionarily doomed animals than we have usually seen. If humans, too, were incapable of escaping evolution's cyclicality, even a dinosaur might generate pathos. In Bell's Before the Dawn, the projected scenes of prehistory are persistently framed as a "cosmic tragedy" (1934: 19) or "comedy" (85) in which, as we have seen, the tyrannosaur is the "hero" (25). In the projection theater humans walk "upon the stage, but as spectators, not actors," passing through the dinosaurs "as if they were air" (23), a "dreamlike" (39) experience that recalls Prospero's fairy masque in The Tempest, which melts "into thin air" like life itself, "such stuff / As dreams are made on" (Shakespeare 2011: 4.1.150, 156-57). Visual access to Mesozoic life prompts sober reflection. One projection technician notes that humanity will one day be just another recorded "spectacle" (24) for unsympathetic future beings, while another proposes that "our overdeveloped central nervous system" is "as outdated as the over-protected, over-armored monstrosities we saw in some of the later records"--although he also speculates, in a more Stopesean mood, that human "psychic development" may be able to "subdue nature" (139). In a similar vein, Jacquetta Hawkes (1951: 35) wondered if the gigantism of the human brain was any less an evolutionary dead-end than "the horn of Synthetoceras." If not, the specialized development of useless "protuberances" could be seen either as a group's last "heroic efforts" (Tolmachoff 1928: 1133), or as an "inner deficiency" (1140) that nonetheless resulted in "high perfection in certain features." Noting the "moral preoccupations" that led scientists to promote one evolutionary mechanism over another, the coauthors of The Science of Life found "something heroic in the obstinate advance of Orthogenesis" (Wells, Wells, and Huxley 1931: 265). Rather than imperial Roman languor, this was a dignified fall caused by tragic hamartia.

These animals faced what Swinton (1948: 219) called "the problem of Hamlet": '"to be or not to be'--specialized." We have already heard some of Swinton's normative pronouncements on the "old and effete" genera described in The Dinosaurs (1934: 179), but the book's elegiac conclusion strayed beyond his characteristic grotesque register. The dinosaurs' "drama," he declared, "was a well-acted one and had an exceptionally long run," tempting Swinton "to anthropomorphize" and wonder if these animals believed themselves "the emperors for all time" (208). He characterized the fossil record as a theater pasted with "the fragments of old play-bills," leaving readers to ponder "what lesson there is" in the final "failure," despite a "long run," of the dinosaurs' great drama (209). Elsewhere, Swinton was more explicit about what the box office decline of spectacular genera meant. In Giants Past and Present, he gloomily predicted that, in the future, "the incurable inhumanity of man to man will have erased the weak" (Swinton 1966: 183). Rather than framing this as a victorious elimination of the unfit in the style of Love's Creation, however, he saw it as a sad victory of "Demos, the god of mediocrity." Based on the paleontological evidence that "it was never the brilliant, the exotic, the spectacular that survived," but rather "the mediocre," he predicted that the future belonged to "small men." For all his mockery of spinescent genera, then, Swinton suspected that long-term evolutionary victory required bland mediocrity.

Although dinosaurs were by then on the verge of being understood as examples of evolutionary success (Ostrom 1969), Swinton's revised 1970 edition of The Dinosaurs differed relatively little from its 1934 original. His overall estimation of the dinosaurs was indicated by an epigraph in heroic couplets:

 They built no cities, shaped no great empires Knew naught of wisdom, had but few desires. They lived each day as life itself compelled, Marked no tomorrow for all it might have held. Friendless in life and all alone in death They left but dust--yet men have given it breath. (Swinton 1970: n.p.)

By this point, however, working at the Royal Ontario Museum and the University of Toronto, Swinton acknowledged that in the postwar decades directed evolutionism had become marginalized, inserting qualifications to several orthogenetic claims (fig. 3). Meanwhile, far wider audiences would still have been reading his more affordable and near-identically titled Dinosaurs, which cast no doubt on "phylogeronty, the old age of a phylum" (Swinton 1962: 36). As Swinton was informed in 1977, a barely altered edition of the 1962 Dinosaurs was still selling well in the London Natural History Museum's bookshop (Cross 1977). Meanwhile, paleontologist Stephen Jay Gould (1973: 16) was informing general readers of Natural History, the popular magazine of the American Museum of Natural History, that the case for orthogenesis had "never rested on more than subjective wonderment," and by then historians were even reconsidering whether what had been called the Fall of Rome was really a fall in the first place (Bowersock 1996: 35; Malik 2019: 43). Indeed, among them the concept of decadence itself had become deeply unfashionable, Harry Ritter's Dictionary of Concepts in History (1986) reporting that theories of decadence were lately deemed overly "value-laden" (99) while "the analogy of the natural life cycle as a guide for scholarship is widely mistrusted" (102). More recently, Neville Morley (2004: 575) attributed the decline of the idea of decadence to its "too obviously literary" nature, hinting, however, that its unpopularity among historians stemmed less from conceptual rejection than from its status as an insufficiently "dead" metaphor.

Conclusion

In William Hope Hodgson's apocalyptic romance The Night Land (1912), evolution comes full circle. In the far future, at a time when humans struggle to survive following the death of the sun, "olden Monsters" resembling those that existed "in the Early World" are "bred in the Ending" (129). For Hodgson, dinosaurs return when we approach the end, like shadows cast by our own decadence. This is an especially literal expression of directed evolution, an idea that, whether in the hands of popular romancers, didactic reformers, or paleontologists, was a potent source of imaginative and political inspiration between the nineteenth-century fm de siecle and the postwar decades. Although it burgeoned at the height of imperialism, and of both the decadent and the eugenic movements, the moralization of orthogenesis and phylogeronty was far from internally consistent, reflecting the tension between those who saw humans as able to combat the process of decay and those who did not. For this reason, directed evolution attracted diverse political constituencies, although, as in many of the texts I've discussed, it was often tied to conservative anxieties about masculine virility as well as racial and class hygiene.

In parallel to such anxieties, running through many of the works that engaged directed evolution, was the fear that overspecialization created weak, alienated individuals and societies. But in addition to signifying harmfully rigid evolutionary trends, overspecialization could also mean the extreme subdivision of labor or of scholarly disciplinarity, and indeed often dimly included all three. The likes of Osborn and Burroughs preached the value of a strenuous life and varied interests as the way to avoid becoming the human equivalent of a specialized, spinescent Triceratops. A very different thinker, Virginia Woolf, also looked to prehistory to challenge the constrictive structures of modern society. Between the Acts (1941), for example, is suffused with imagery of anatomically generalized prehistoric animals, as Evelyn Tsz Yan Chan (2014: 154) points out, arguing that it enacts a devolutionary "retreat from overspecialization" to help "reach a balance in modern life." Orthogeneticists, of course, would have warned that prehistory itself contains countless precedents for overspecialization as well, but Woolf, a reader of Wells serials like The Science of Life, had little time for teleological interpretations of life's development. Catriona Livingstone (2022: 194) adds that in Between the Acts and other late Woolf novels, "sympathetic identification with other members of the community," including animals, "causes characters to become aware of and identify with unrealized aspects of their selves," questioning the social and evolutionary grooves down which they are being unconsciously driven. Rather than structuring evolution and social history as a deterministic series of rises and declines, Woolf asserted the emancipatory potential of overlaps and unexpected affinities.

As an alternative to Woolf's modernistic vision, directed evolution's appeal was that it offered in decadence a linear but resonant narrative structure applicable to the prehistoric, ancient, and modern worlds alike, along with a new opportunity to take charge of historical progress. Its potential fatalism gave rise to the contradictions I have highlighted: authors looked on phylogerontic animals mostly with scorn, but sometimes with empathy and awe. Repeatedly, these animals were cited as evidence in favor of prioritizing flexibility over habit, brains over brawn--but not without the sense that something important and even wonderful was being lost along the way. Even before paleontologists began more regularly to consider structures attributed to orthogenesis as adaptive products of natural and sexual selection, these reflections were sobering. Hinting that decadence might in some ways be valuable, they imply that there are criteria for determining value other than resilience, moderation, and even intelligence. In this light, the forked horn of Synthetoceras did not look quite so ridiculous.

DOI 10.1215/0041462X-11098327

Richard Fallon is a postdoctoral researcher at the University of Nottingham and the Natural History Museum, London. He is the author of Reimagining Dinosaurs in Late Victorian and Edwardian Literature: How the "Terrible Lizard" Became a Transatlantic Cultural Icon (2021).

Acknowledgments

This work was supported by the Leverhulme Trust (grant ECF-2020-055).

Notes

(1.) Pinner's vivid illustration, and Swinton's accompanying text, were likely the inspiration for Jacquetta Hawkes's selection of this obscure genus as her example of the perils of directed evolution.

(2.) These descriptions recall the orgiastic and opulent paintings of John Martin, especially Belshazzar's Feast (ca. 1821) and Pandemonium (1841).

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Caption: Figure 1: Erna Pinner, "Restoration of the Extinct Deer Synthetoceras Illustration for The Corridor of Life, by William Elgin Swinton (1948). Reproduced with the kind permission of Frances Kitson, Jacky Oldham, and Peter Oldham.

Caption: Figure 2: A diagrammatic demonstration of the analogy between spiny skin on an elderly individual animal and in an "elderly" species (Beecher 1898: 355). Reproduced courtesy of the Cadbury Research Library, University of Birmingham.

Caption: Figure 3: William Elgin Swinton inserts qualifications to his earlier comments on the "decadent" status of the later dinosaurs in the 1970 revision of his 1934 work, The Dinosaurs (Swinton n.d.). Courtesy of the Royal Ontario Museum.